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BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA:
                                  AN UPDATE
     A REPORT BY DAVID PEACOCK, TARNYA COX, TANJA STRIVE, GREG MUTZE,
                                     PETER WEST AND GLEN SAUNDERS

COLLABORATION              INNOVATION                   IMPACT
BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: Centre ...
In memory of Greg Mutze 1957-2021

Published by:

Centre for Invasive Species Solutions

Building 22
University Drive South
University of Canberra
Bruce, ACT 2617

Mailing Address:

PO Box 5005,
University of Canberra LPO,
University of Canberra ACT 2617
Telephone: 02 6201 2887
Home Page: www.invasives.com.au

ISBN Print: 978-1-925727-35-7

ISBN Web: 978-1-925727-34-0

Acknowledgement:

The Centre for Invasive Species Solutions gratefully acknowledges the significant contributions and input into this report
from a range of stakeholders and experts, in particular David Peacock and Ken Moore as well as Tarnya Cox, Tanja Strive,
Greg Mutze, Peter West, and Glen Saunders, who were authors of the first Benefits of Rabbit Biocontrol report in 2013
and whose publication forms the foundation on which this update is based.

Citation:

Centre for Invasive Species Solutions (2021). Benefits of Rabbit Biocontrol in Australia: an update. Centre for Invasive Species
Solutions, Canberra. David Peacock, Tarnya Cox, Tanja Strive, Greg Mutze, Peter West and Glen Saunders.
December 2021.

Copyright:

This publication is copyright. Apart from fair dealing for the purposes of private study, research, criticism or review as
permitted under the Copyright Act 1968, no part may be reproduced, copied, transmitted in any form or by any means
(electronic, mechanical or graphic) without the prior written permission of the Centre for Invasive Species Solutions.

This report may be cited for purposes of research, discussion, record keeping, educational use or other public benefit,
provided that any such citation acknowledges the Centre for Invasive Species Solutions.

Disclaimer:

While great care has been taken in ensuring that the information in this report is factual and accurate at the time of
publication, however the Centre for Invasive Species Solutions accepts no responsibility for omissions, errors, or changes
in data since the time of publication.

Cover Images:

Native orchid regeneration after RHDV reduced rabbit grazing, Coorong National Park (SA), Sept. 2016 (photo by
David Peacock). Rabbit image by Laurence Sanders.

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Contents

TABLE OF CONTENTS
List of Figures                                                                     4

List of Tables                                                                      4

Executive Summary                                                                   5
Glossary of abbreviations                                                           7

Introduction                                                                       8
Overview of rabbit distribution, abundance and impacts                             8
Environmental impacts                                                              10

Examples of local environmental impacts from browsing                              11

Impacts on native animals                                                          13

Economic impacts                                                                   14

Social impacts                                                                     15

History and Benefits of Rabbit Biocontrol in Australia                             16
Environmental benefits                                                             21

The recovery of native vegetation                                                  21

Native mammal recovery                                                             26

Biosequestration                                                                   28

Economic benefits                                                                  28

Social benefits                                                                    29

A holistic picture of the benefits of rabbit control                               29

Need for future rabbit biocontrol agents                                           30
References                                                                         32
Appendix 1: Background Information on Rabbit Biocontrol Agents                     38
Myxomatosis                                                                        38

Rabbit fleas                                                                       38

Rabbit haemorrhagic disease                                                        38

Virally-vectored immunocontraception                                               39

Eimeria parasites (an option recommended by the Rabbit Biocontrol Business Case)   40

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LIST OF FIGURES
Figure 1: Rabbit ancestry genetics in Australia.                                                                            8

Figure 2: Rate of spread of wild rabbits since their primary introductions in 1857–60 to their known occurrence as reported
          in 2012.                                                                                                          9

Figure 3: Reported abundance of rabbits across Australia.                                                                  9

Figure 4: Locations of critically endangered and endangered communities and species that may be adversely affected by
          rabbits.                                                                                                    10

Figure 5: Rabbit grazing of the understory and browsing of this dryland tea tree (Melaleuca lancelota) to rabbit height.   11

Figure 6: Ring barking of large native pines by rabbits.                                                                   12

Figure 7: Native plants that are susceptible to rabbit browsing.                                                           13

Figure 8: Illustration of general trends in rabbit numbers since the release of myxomatosis (1950) and likely benefits
          of subsequent releases of biocontrol agents into the Australian rabbit population.                               17

Figure 9: Rabbit biocontrol innovation pipeline.                                                                           20

Figure 10: Average decline in rabbit populations by rainfall zone after the arrival of RHDV1.                              21

Figure 11: Native vegetation regeneration on Thackaringa Station, Broken Hill.                                             22

Figure 12: Recovery of arid shrublands in northern South Australia as detected by Landsat satellites.                      23

Figure 13: Regeneration of native species at Turretfield, South Australia, October 2017. post-RHDV2.                       24

Figure 14: Increase in the native orchard population in the Coorong National Park following the establishment of RHDV1
           and the suppression of rabbit numbers.                                                                          25

Figure 15: Landscape scale recovery of threatened desert mammals following the release of RHDV1.                           26

Figure 16: Landscape scale suppression of rabbits, foxes and feral cats at Roxby Downs, South Australia, in 1989-90
           following arrival of RHDV1.                                                                                     27

Figure 17: How the benefits of control of rabbits are integrated.                                                          30

LIST OF TABLES
Table 1: A historical timeline of rabbit biocontrols in Australia.                                                         16

Table 2: Estimated present value of benefits for RHDV1-K5 under different scenarios regarding the presence of RHDV2.
Source: Hardaker and Chudleigh (2020).                                                                                     29

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EXECUTIVE SUMMARY
Australians have been battling the impacts of the European rabbit (Oryctolagus cuniculus) since the mid-19th century .

This report provides the evidence base for ongoing investment in biocontrol research, development and engagement
(RD&E), and the release of new agents to further suppress rabbit abundance when appropriate. It should be read in
conjunction with the Centre’s updated Rabbit Biocontrol Pipeline Strategy (CISS 2021).

This report also documents key parts of the substantial body of research that analyse the economic, environmental and
social impacts of rabbits on Australia, the effectiveness of biocontrol, and qualitative and quantitative benefits that derive
from biocontrol.

Historical context
Following its successful introduction on mainland Australia between 1857 and 1860, the European rabbit became the fastest
colonising terrestrial mammal anywhere in the world due to a lack of natural predators and its successful adaption to a
variety of environments. The rapid expansion of rabbits across Australia was enhanced by their prolific breeding potential.
Rabbits can begin reproducing at four months of age and, in favourable conditions, may produce five or more litters per year
that results in between 50 to 60 offspring per year per female rabbit. Under favourable conditions, the reproductive activity
of a single pair of rabbits can result in a population of 184 rabbits in 18 months. Within 70 years of their release, rabbits had
spread to cover almost 70 per cent of Australia’s landmass, to an area estimated at 5.3 million square kilometres.

Rabbits quickly became Australia’s worst terrestrial vertebrate pest with plagues causing major damage to agriculture, the
developing economy, rural communities and the environment. The National Museum of Australia has identified the success
of their introduction to the continent as one of the “defining moments in Australian history”.

Initial control efforts centred around trapping, rabbit warren ripping, fumigation and bounty systems. Later, as noted by the
Museum, “Fences became an integral component of what settlers in the late 19th century began to see as a war against the
rabbits.” It is estimated that there were 320 000 kilometres of rabbit proof-fence across Australia at the height of the fencing
boom. Colonial governments also moved to make it compulsory for landholders to destroy all rabbits on their farms and
stations and made it illegal to aid their spread. Despite these early rabbit control efforts, Australians were fighting a losing
battle.

In 1943, in response to growing public pressure to find alternative rabbit control solutions, CSIRO conducted initial tests
using the rabbit specific myxoma virus (MV). These tests initially appeared to fail but following good rain in late 1950 the
virus was rapidly spread by mosquitos. It had a dramatic effect on the rabbit population, then numbering around 600 million,
reducing rabbit numbers by as much as 99 per cent especially in arid areas. Infections were further assisted by the release
of two flea vectors in 1969 and 1993.

The major success of the myxoma virus as a biocontrol put Australia on the path to becoming world leaders in the
management of pest rabbits. However, as has been the experience of later biocontrol agents, resistance to MV in the rabbit
population led to a gradual recovery in their numbers.

The continued development of biocontrol knowledge for controlling rabbits through RD&E has become essential in the
ongoing fight against wild rabbits. It has enabled the 1995 release of the rabbit haemorrhagic disease virus, RHDV1, and a
new strain, RHDV1-K5, in 2017. The release of RHDV1 initially reduced the rabbit population by up to 98 per cent, particularly
in arid areas.

Serendipitously, a new viral agent, RHDV2, appeared in Australia from unknown sources in 2014. In less than two years
RHDV2 had spread through the Australian rabbit population. This had a significant initial impact and reduced rabbit
abundance by 60 per cent on average. However, its emergence also restricted the ability of RHDV1-K5 to spread within rabbit
populations. While this led to the impacts of the managed release of RHDV1-K5 being suppressed, combined they effectively
impacted the rabbit population. At present RHDV1 and RHDV2 are coexisting in the environment however it is uncertain if
this will be the case into the future. RHDV2 may become the dominant field strain.

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The impact of rabbits and the benefits of rabbit biocontrol
Prior to the release of the myxoma virus, the financial impact of rabbits on agriculture was some $2 billion a year. Rabbit
biocontrol based on the myxoma virus and RHDV1 Czech 351 strain reduced this impact to about 15% of the pre-release
cost. This is equivalent to $81.8 billion in agricultural productivity in the sixty years to 2011 (expressed in 2020-dollar terms;
Reserve Bank of Australia, 2022). The RHD viruses introduced (or naturalised) since 2014 — that is, RHDV2, RHDVa and
RHDV1 K5 — are expected to generate benefits of a further $4 billion over the next 30 years. This is on top of the ongoing
benefits derived from myxoma virus and the RHDV1 Czech 351 strain.

Rabbit biocontrol has had major benefits for native plant and animal species, and in arid inland Australia has been
attributed as the single most important and cost-effective conservation action for small, threatened mammals and a range
of ecosystems in recent decades. An associated benefit enabling their recovery has been a reduction in abundance of foxes
and feral cats which is directly attributed to the removal of their primary prey, the rabbit.

While the benefits from rabbit control have been enormous, rabbits persist in the Australian landscape and continue to
have a severe impact. Many Australian native plants are especially sensitive to rabbit browsing. Recruitment for some slow
growing species such as Mulga are negatively impacted by rabbits with densities as low as one rabbit/km2 (0.01 rabbit/ha).
The impact rabbits have on both native plants and animals is the reason they impact 322 nationally threatened species – the
most for any vertebrate pest. They also continue to cause $217 million a year in lost agricultural productivity.

In summary, controlling rabbits at a landscape scale with biocontrol methods results in less erosion and weed growth and
fewer feral predators of native animals. Importantly, the use of biocontrol is critical to reducing the abundance of rabbits
at large scales to less than 0.5 rabbits per hectare allowing the survival and growth of palatable native plant seedlings.
This leads to more native animals, plant species and vegetative growth. It also increases carbon sequestration, makes
food production more sustainable, and improves ecosystem and landscape health — factors essential to the welfare and
prosperity of all Australians.

While a silver bullet for the eradication of wild rabbits remains elusive, biological control over the vast range of rabbit
distribution in Australia has been highly successful. However, biocontrol remains an ongoing process: rabbit populations
continue to have the ability to build resistance to introduced pathogens and for their abundance to increase.

The need for ongoing biocontrol RD&E has been recognised by governments, research agencies and industry organisations
and has led to two Centre five-year R&D plans under a long-term Rabbit Biocontrol Pipeline Strategy:

      — Phase two of the Strategy was implemented by the Invasive Animals Cooperative Research Centre (2012–2017)

      — Phase three of the Strategy is currently being implemented by the Centre for Invasive Species Solutions
        (2017–2022).

      — Phase four of the Strategy is currently being prepared by the Centre for Invasive Species Solutions (2022–2027).

Phase four offers the potential of new biocontrols that include genetic biocontrol technologies and the monitoring of rabbit
abundance impacts through new satellite imaging methodologies and the use of artificial intelligence.

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Glossary of abbreviations
CSIRO   Commonwealth Scientific and Industrial Research organisation.

IA CRC Invasive Animals Cooperative Research Centre that has transitioned into the Centre for Invasive Species Solutions.

MV      Myxoma Virus, the first successful rabbit biocontrol intentionally released in Australia in 1950.

RCV-A1 An endemic benign Rabbit Calicivirus detected in 2009 and which provides partial protection to lethal RHDV1
       infection.

RHDV(s) Rabbit Haemorrhagic Disease Virus ­— used in this report as a generic descriptor for all Rabbit Haemorrhagic
        Disease viruses.

RHDV1 The initial Czech-351 strain of Rabbit Haemorrhagic Disease Virus released in Australia in 1995

RHDV1-K5 The naturally occurring Korean strain of Rabbit Haemorrhagic Disease Virus released in Australia in 2017

RHDV2 The variant of RHDV detected in Australia in 2014 that was not intentionally imported or released.

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BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: Centre ...
INTRODUCTION
This report reviews the distribution, abundance and impacts — environmental, economic and social — of European rabbits
(Oryctolagus cuniculus) since the mid-19th century to the present day. It examines the history and effectiveness of the
biocontrol agents that were released in Australia — MV, RHDV1, RHDV1-K5 and two flea vectors — or serendipitously
appeared (RHDV2). It outlines the demonstrated environmental, economic and social benefits of rabbit biocontrols in
Australia, and makes the demonstrable case for ongoing national investment in rabbit biocontrol research, development
and engagement (RD&E).

This report should be read in conjunction with the Centre’s report on the third phase of the Rabbit Biocontrol Pipeline
Strategy (CISS 2021).

Overview of rabbit distribution, abundance and impacts
Wild rabbits were successfully introduced onto the Australian mainland in the mid to late 19th century at multiple locations,
with primary releases in South Australia and Victoria between 1857 and 1860 (Peacock and Abbott 2013). An analysis of
rabbit ancestry genetics suggests there were a number of successful rabbit introductions to Australia (see Figure 1; Iannella
et al. 2019).

Within 70 years rabbits had spread from release locations to inhabit 70 per cent of Australia’s landmass (5.3 million km2).
They are widespread throughout most locations where they are found (NLWRA and IA CRC 2008). Figure 2 shows the rate of
spread of wild rabbits since their primary introductions in 1857-60 to their known distribution as reported in 2012. Figure 3
shows the reported abundance of rabbits across Australia including rabbit sightings as recorded in RabbitScan from 2009 to
2013.

        Figure 1: Rabbit ancestry genetics in Australia. Pie charts indicated proportion of ancestry at each site as estimated by fastSTRUCTURE with
        a K=6. Historical rabbit introduction records of successful or unknown outcome reported by Peacock and Abbott (2013) are represented
        as red diamonds, noting some may be hidden by pie charts. The Barwon Park release site in Victoria and state capital cities are specifically
        indicated. Red diamonds indicate historical introduction points (Ianella et al. 2019).

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Figure 2: Rate of spread of wild rabbits since their primary introductions in 1857–60 and their known occurrence as reported
          in 2012. (IA CRC. Stodart amd Parer. 1988).

       Figure 3: Reported abundance of rabbits across Australia (IA CRC, NLWRA/IA CRC 2008, RabbitScan 2009–13).
BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: Centre ...
Whether rabbits have reached the limits of their absolute range is not certain. The ‘Background document to the Threat
abatement plan for competition and land degradation by rabbits’ (DAWE, 2016a) cites the view of NLWRA and IA CRC (2008)
that rabbits have largely reached their ecological limit in terms of range. Since rabbit populations respond to weather
conditions, they are likely to extend their range into areas they currently do not occupy when these are favourable but when
weather conditions deteriorate, their range may contract back to previous limits. As such, the average range of rabbits may
have been reached, although climate change predictions for hotter temperatures could reduce their average range over
coming decades. For example, Pavey and Bastin (2014) predict that rabbits could become absent from large areas of the
centre and west of the NSW Rangelands by 2050.

The rapid expansion of rabbits across Australia was enhanced by their prolific breeding potential. Rabbits can begin
reproducing at four months of age and, in favourable conditions, may produce five or more litters per year (NSW DPI 2007).
A short gestation period coupled with females generally mating within hours of giving birth can result in between 50 to 60
offspring per year per female rabbit. Under favourable conditions, the reproductive activity of a single pair of rabbits can
result in a population of 184 rabbits in 18 months (Agriculture Victoria 2021).

Environmental impacts
When rabbits feed they remove vegetation to ground level, browsing, chewing bark and ringbarking, and digging for roots.
This can result in significant soil destabilisation and erosion, leading to the desertification of heavily impacted areas. Their
excavation of extensive warrens adds to this damage, particularly in ecological sensitive areas. This is further exacerbated
by their prolific breeding — in good seasons their populations can increase seven to tenfold. Overall, rabbits have an impact
on 322 nationally threatened species (Kearney et. al. 2018) and nine ecological communities listed under the Commonwealth
Environment Protection and Biodiversity Conservation Act 1999 (DAWE 2016b) (Figure 4).

        Figure 4: Locations of critically endangered and endangered communities and species that may be adversely affected by
                  rabbits.

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Research by the former Invasive Animals Cooperative Research Centre (IA CRC) and its partners has shown that Australian
native vegetation is very sensitive to rabbit damage. A specific example is mulga (Acacia aneura), which occurs across vast
areas of central Australia. Recruitment of mulga seedlings is negatively impacted by rabbit densities as low as one rabbit/
km2 (0.01 rabbits/ha) (Mutze et al. 2008). This greatly impedes the natural regeneration of mulga in many areas where
rabbits are present in central Australia. Rabbits have similar negative impacts on the recruitment of a variety of palatable
trees and shrubs across much of their distribution in Australia (Cooke 2012, Forsyth et al. 2015, Mutze et al. 2016a). Highly
palatable species are severely damaged from selective grazing pressure at low densities equal or greater than 0.5 rabbits
per hectare, with moderately palatable plant species being severely damaged at two rabbits per hectare (Mutze et al 2016b).

Examples of local environmental impacts from browsing

Figure 5 shows a dryland tea tree (Melaleuca lancelota) that grew during a period of rabbit relief after the spread of MV,
but with increasing rabbit numbers due to developing resistance to MV, the understory has been removed by grazing.
The photograph also shows browsing of the tree to rabbit height.

        Figure 5: Rabbit grazing of the understory and browsing of this dryland tea tree (Melaleuca lancelota) to rabbit height (Photo: Brian Cooke).

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Figure 6 shows ring barking of large native pines (Callitris glaucophylla) by rabbits at Holowiliena, South Australia, during winter 1991 and pre-RHDV1 release.

                          Figure 6: Ring barking of large native pines by rabbits. (Photo: David and Carol Warwick)

        Many other native species are susceptible to rabbit browsing. Figure 7 shows browsing occurs even at low
        rabbit densities and during periods when other feed is available which is detrimental to both the growing
        plant and the level of recruitment to those plant populations.

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Figure 7: Native plants that are susceptible to rabbit browsing: Top left: Boobiala (Myporum insulare), Coorong SA. Top right: Sandhill wattle (Acacia ligulate),
Hattah-Kulkyne National Park, VIC. Bottom left: White cypress pine (Callitris glaucophylla), Ikara-Flinders Ranges National Park, SA. Bottom right: Buloke
(Allocasuarina luehmannii), Hattah-Kulkyne National Park, VIC. (Photos: Brian Cooke & David Peacock).

Impacts on native animals
While the direct impact of rabbits on vegetation has been clearly documented, their impact on native animals via competition
is less clear. Both kangaroo (Macropus rufus, M. fuliginosus, and M. robustus) and wombat (Vombatus ursinus) populations
increased following native vegetation regeneration after the release of RHDV and the sharp decline in rabbit abundance. This
suggests that high rabbit populations may restrict these species (Mutze et al. 2008; Bird et al. 2012). A similar explanation
has been proposed for the very restricted mainland distribution of the quokka (Setonix brachyurus) (Scholtz and DeSantis
2020).

The impact of rabbits on native pastures directly affects a variety of other species, including endangered or critically
endangered species such as the plains-wanderer (Pedionomus torquatus) (Baker-Gabb 2002) and the golden sun moth
(Synemon plana) (Clarke and O’Dwyer 2000). Competition and land degradation caused by rabbits is listed as a ‘Key
Threatening Process’ under the Commonwealth Environmental Protection and Biodiversity Conservation Act 1999.

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Economic impacts
Rabbits have had a major economic impact on Australian agriculture from the late 19th century to the present day. In
earlier times, when agriculture was a major part of the nation’s gross domestic product, the scale of this impact was felt
on the national economy. Prior to the release of the myxoma virus, the financial impact of rabbits on agriculture was
some $2 billion a year. (Cox et al. 20130).

There have been many studies of the economic impact of rabbits on agriculture especially for wool, sheepmeat and beef
production. The most contemporary comprehensive studies have been McLeod (2004), Gong et al. (2009) and McLeod et al.
(2016). Other significant economic studies such as Cooke et al. (2013) and Hardaker and Chudleigh (2020) have focused on
quantifying the economic benefits of biocontrol and are discussed later in this report.

The economic cost of rabbits is most pronounced in Australia’s pastoral industries, however they remain significant pests
in cropping, viticulture and horticulture. The main impacts on livestock industries are overgrazing of native grasslands,
native pastures and improved pastures, a loss of plant biodiversity in pastures, and reduced livestock carrying capacity. The
Department of Primary Industries and Fisheries Queensland (2008) have estimated that rabbits eat around fifteen per cent
of their body weight per day — an amount five times greater than sheep and cattle which consume three per cent of their
body weight per day.

The most comprehensive study on the economic impacts of vertebrate pests in Australia, including rabbits, is by Gong et al.
(2009). This study quantified the direct economic impacts of invasive animals on agriculture (beef, wool, lamb and grains)
in Australia, and the nationwide expenditure by governments and landholders on pest management, administration and
research.

The economic impact from rabbits that Gong et al. identified was grazing competition. This saw a reduction in the carrying
capacity of farmland. The grazing competition resulted in less livestock being carried, lower wool production per animal,
reduced lambing percentage, lessoned wool quality, reduced sale weights and higher stock mortality.

Gong et al. calculated the overall impact of vertebrate pests as the sum of the effects on agriculture plus the expenditure on
their management. Using the concept of the impact on economic surplus, Gong et al. estimated the total annual loss from
rabbits was $206 million.

McLeod (2016) provided an update from the 2009 study of Australia-wide annual production loss costs and expenditure by
governments and landholders on pest animal management including rabbits for 2013 –14. Assuming a fixed product price,
McLeod estimated the annual production loss impact of rabbits for wool, sheepmeat and beef production at $217 million
in 2013 –14 for an average rabbit impact scenario. McLeod also estimated production losses for low and high rabbit impact
scenarios at $108 million and $251 million.

Care should be taken in generalising the results from studies such as Gong et al. (2009) and McLeod (2016) as an estimate of
the ongoing annual cost for agriculture. The results are based on a large number of assumptions for which there are limited
data — or the data changes from year to year — including product prices.

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Social impacts
There has been little systematic research into the social impacts for any vertebrate pests in Australia with the exception of
wild dogs (Fitzgerald and Wilkinson 2009, Thompson et al. 2013). However, there are some studies that look at the impacts of
rabbits on people and communities. These particularly relate to the rabbit plagues of the late 19th and early 20th centuries.

Cooke (2017) examined the impact of rabbits on the economy and culture of the Diyari people of north-eastern South
Australia. He argued that the catastrophic impacts of rabbits on their desert habitats and food resources was the primary
factor leading the Diyari people being driven from their lands and becoming reliant on European support.

The impacts on pastoralists were also severe. There are accounts of pastoralists continuously battling rabbits and then
having to abandon their pastoral leases because rabbits had eaten everything (Rolls 1969, Coman 1999, Cooke 2017). Similar
accounts were given to the associated commissions and royal commissions during the 1890s (Anon 1890, 1893, 1898).

Quealy (2011) provides a collection of personal stories from those impacted by rabbits who were forced to leave their
properties prior to the introduction of the myxoma virus. Many of these speak of the psychological distress at having to
make a new life away from the farm.

The psychological distress and changing attitudes towards rabbits experienced by those battling these plagues were
recorded by early CSIRO scientist, Francis Ratcliffe (Ratcliffe 1938). Such distress was sometimes exacerbated by conflict
in communities arising between those prospering from high rabbit numbers such as commercial harvesters and pest
controllers, and those suffering because of rabbit plagues (Coman 1999). This is perhaps best illustrated by the failed
introduction of the mongoose to Australia as a tool to help control the plagues of the late 1880s, with the rabbiters that
were making their living from killing rabbits being reported as a primary reason for the demise of the mongoose (Peacock
& Abbott 2010).

More recently a study undertaken in the Hunter Valley region of NSW highlighted that residents were concerned by the
impact of rabbits on mine rehabilitation sites, damage to grape vines and the risk of injury to horses due the presence of
warrens (Fitzgerald and Wilkinson 2009).

The National Museum of Australia has acknowledged the social and other impacts of rabbits on Australia since their
successful introduction to the continent as a “defining moment in Australian history” (National Museum of Australia 2021).

The Museum discusses the different methods of rabbit control that were applied and how: “Fences became an integral
component of what settlers in the late 19th century began to see as a war against the rabbits.” Fencing was government
sponsored and at the height of the fence construction boom there were 320,000 kilometres of rabbit proof-fence across
Australia. The longest spanned 3,256 kilometres north to south across Western Australia.

As part of the rabbit fencing boom, colonial parliaments passed legislation to make it compulsory for landholders to destroy
all rabbits on their farms and stations and made it illegal to aid the spread of rabbit.

The Museum notes that fencing failed for many reasons and by the late 1940s the rabbit population had increased to
600 million due to a number of high rainfall years but also because WWII reduced manpower for trapping and fence
maintenance.

At the same time as the country was fighting rabbit plagues, families benefited from the availability of rabbits as a free
source of meat during the Great Depression and itinerant ‘rabbitohs’ or rabbiters found employment in the capture and sale
of rabbits.

In the more prosperous years following WWII, the consumption of rabbits and employment of of rabbitohs largely
disappeared. A small meat rabbit breeding industry developed mainly to source restaurant demand based on New Zealand
white rabbits due to their high feed to meat ratio and fine bones.

There is also a small pet industry for rabbits in some states. Queensland has legislation which makes the keeping of a rabbit
as a domestic pet illegal. In states where rabbits can kept as pets, breeds attractive to children are largely used.

The existence of the small meat and pet rabbit industries mean that introduced biocontrols for wild rabbits need to ensure
protections for farmed and pet rabbits, largely through the availability of vaccines.

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HISTORY AND BENEFITS OF RABBIT BIOCONTROL
IN AUSTRALIA
Australia has introduced four biocontrol agents for rabbits over the past 60 years: two viral diseases (myxoma virus – MV
and rabbit haemorrhagic disease virus – RHDV1) and two flea vectors (Spilopsyllus cuniculi and Xenophsylla cunicularis) to
aid their transmission. A third viral agent, RHDV2, was detected in Australia in 2014, but it is not known how the virus entered
the country. A new strain of RHDV1, K5, was released nationally in 2017. A detailed timeline showing the history
of biocontrols is shown in Table 1.

Biological control agents have been the most successful method of reducing rabbit abundance in Australia by far. The initial
impact of the MV reduced rabbit numbers by as much as 99 per cent and RHDV1 by up to 98 per cent in arid areas. The
general trends in rabbit numbers since 1945 and projected numbers through to 2029 and beyond are illustrated in Figure 8.

The general trend across Australia following the release of a new biocontrol or vector has been a rapid and substantial
decline in rabbit abundance, followed by a gradual recovery in numbers. Despite such increases, rabbit abundance has
generally not reached the pre-biocontrol release levels.
Table 1: A historical timeline of rabbit biocontrols in Australia.

 1857–60                            The primary rabbit releases at Barwon Park (Vic), Anlaby Station (SA) and Point Lowly (SA) for hunting become
                                    established. Some farms abandoned as early as 1881.
 Late 1940s                         The rabbit population increases to 600 million due to a number of high rainfall years and because of reduced
                                    manpower for trapping and fence maintenance during WWII.1
 1950                               The world’s first vertebrate pest biocontrol — myxoma virus (MV) — released, killing 99.8% of infected
                                    rabbits.
 1951–52                            Rapid selection of genetically resistant rabbits leads to weakened forms of myxomatosis.
 1969                               Rabbit flea species (Spilopsyllus cuniculi) approved for release to act as an improved vector to spread MV in
                                    areas with low mosquito levels.
 1993                               Spanish rabbit flea (Xenophsylla cunicularis) adapted for arid conditions approved for release and improves
                                    transmission of MV.
 1995                               The rabbit population returns to about 300 million with populations continuing to rise.2
 1995                               Rabbit haemorrhagic disease virus (RHDV1) introduced, killing up to 98% of rabbits in arid areas.
 2000s                              Rabbits begin to develop resistance to RHDV1 infection. Increasing resistance results in increasing rabbit
                                    numbers being observed.
 2009                               Benign endemic rabbit calicivirus (RCV-A1) discovered and characterised.3 RCV-A1 confers partial protection
                                    to lethal RHDV1 infection and therefore impedes effective RHDV based biocontrol.
 2009+                              To counteract RCV-A1, new naturally occurring RHDV1 strains from overseas are imported from Europe and
                                    Asia and evaluated as part of the RHD Boost project to increase the effectiveness of RHDV1 based biocontrol.
 2012                               Invasive Animals CRC develops 20-year rabbit biocontrol pipeline strategy and strategic rabbit biocontrol
                                    research program to boost RHDV1 effectiveness, assess feasibility of new potential rabbit biocontrol
                                    candidates, and increased capabilities to promote integrated rabbit control at
                                    a regional level.
 2014                               Rabbit haemorrhagic disease virus 2 (RHDV2) arrives in Australia and spreads to all rabbit populations across
                                    Australia within 2 years. Rabbit populations reduced by 60% on average and up to 80% in some populations,
                                    including a proportion of rabbits with immunity to RHDV.
 2017                               The first nationally coordinated release of a new RHDV1 strain, RHDV1-K5, in 20 years (based on former
                                    Invasive Animals CRC program). Delivers a 34% national average knockdown at release sites but is
                                    outcompeted by RHDV2 at a landscape scale.4
 2016–18                            RHDV2 becomes the dominant strain in the Australian landscape.5
 2017                               Centre for Invasive Species Solutions formed as successor to IA CRC and continues to advance rabbit
                                    research, including the assessment of RHDV2 as a biocontrol agent.
 2018                               Scientists discover that rabbits that survive MV have a 10% poorer survival rate when subsequently infected
                                    with RHDV1. The interaction between biocontrols provides additional benefit.6
 2019                               Intestinal Eimeria rabbit parasites assessed as an additional control agent but found to be unsuitable as
                                    virulent Eimeria parasites are already widespread across Australia.7

 1
  National Museum of Australia (2021). 2Ward (2011), 3Strive et al (2009). 4Cox et al (2019). 5Ramsey et al (2020). 6Barnett et al (2018). 7Peacock et al (under
 review) Source: Adapted from the Victorian Department of Environment and Primary Industries (2011). See Appendix 1 for description of each biocontrol
 technique mentioned above.
This pattern is reflected in rabbit abundance in the arid pastoral area of north-eastern South Australia following with the
introductions of MV, RHDV1, the arrival of RHDV2 and then the release of RHDV1-K5 (Figure 9). The horizontal red line on the
graph shows the relative rabbit abundance above which recruitment of palatable native vegetation does not occur.

Other control measures, such as poisoning and warren ripping were not often applied in this region so population variations
can be primarily attributed to the biocontrol agent. As mosquitoes are generally scarce in the region, the release of European
rabbit fleas (Spilipsyllus cuniculus) in 1969 enhanced the spread of MV (Cooke et al. 2013).

Figure 8: Illustration of general trends in rabbit numbers since the release of myxomatosis (1950) and projected benefits of subsequent releases of biocontrol
agents into the Australian rabbit population. The rabbit impact threshold shown is for illustration purposes only as impact thresholds vary for different
landscapes, ecological communities and species. In reality, rabbit numbers as low as 1 rabbit per 2 ha can have significant detrimental impacts to the
environment. It is important to note that conventional rabbit control is essential to maintaining low rabbit levels and capitalising on the impact of biocontrol
(Modified from Saunders et al 2010, Cox et al. 2013, Cooke 2018, Strive & Cox 2019)

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A LONG-TERM RABBIT BIOCONTROL PIPELINE
STRATEGY
The Centre for Invasive Species Solutions (CISS) — and its predecessor, the Invasive Animals Cooperative Research Centre
(IA CRC) — together with its members and partners, recognise the environmental, economic and social benefits of long-
term sustainable rabbit control. These organisations have championed a multi-pronged, long term strategic approach
to harness the unique opportunities and potential high returns on investment that successful biocontrol initiatives can
provide. This approach is detailed in the Rabbit Biocontrol Pipeline Strategy, 2012–2027 (CISS 2021, in production).
A summary of rabbit biocontrol pipeline activities undertaken since the 1960s is presented in Figure 9.

Phase 1, IA CRC foundational activities 2007 to 2012
The foundational activities of the rabbit biocontrol pipeline strategy were undertaken between 2007 and 2012 by the IA
CRC. The primary focus of these activities was on the RHDV strains already present in Australia and understanding the
geographic variation in their efficacy.

CSIRO research through the IA CRC was the first to uncover the presence of an endemic, non-pathogenic form of
calicivirus — Australian rabbit calicivirus (RCV-A1) (Strive et al 2009) — and showed that it was able to provide transient
and partial immunity to lethal RHDV1 infection (Strive et al 2013). Experimental infection studies showed that while some
rabbit populations were developing genetic resistance to infection with RHDV1, the virus was partially compensating for
this by evolving towards relatively increased virulence.

Following this work, over the past decade, governments and industry have co-invested in two five-year R&D plans, driving
forward the long-term rabbit biocontrol pipeline strategy.

Phase 2, 2012 to 2017
The ‘RHD Boost’ project was conceived when it became apparent that the prevalence of RCV-A1 was impeding the
performance of the initial RHDV1 strain (Czech 351) in cool, high rainfall areas of the continent (Strive et al. 2010; Liu et
al. 2014). RHD Boost aimed to identify, evaluate and select any suitable naturally occurring overseas strains of RHDV that
might be able to outcompete RCV-A1 and increase the effectiveness of RHDV in higher production, temperate areas of
Australia.

Once 38 RHDV variants and RHDV-like viruses had been imported and evaluated by RHD Boost, the Korean K5 strain
(RHDV1-K5) was selected as the biocontrol agent for release. This was based on its increased ability to overcome partial
cross protective immunity provided by the non-pathogenic RCV-A1 and its increased ability to infect genetically resistant
wild rabbits (IA CRC 2014). A complementary study confirmed the efficacy of a common commercial RHDV1 vaccine on
RHDV1 K5 (Read and Kirkland 2017).

Approval for the use of RHDV1-K5 was given be the Australian Pesticides and Veterinary Medicines Authority (APVMA)
and other government regulators prior to its nationwide release in March 2017. Before its release, increased surveillance
efforts put in place by RHDV Boost identified the exotic RHDV2 in Australia in 2015, and later demonstrated it had arrived
earlier in 2014. RHDV2 spread rapidly within the Australian rabbit population — within 18 months it had mostly replaced
the existing RHDV1 strain and become the dominant calicivirus in the Australian landscape. The impact of RHDV2 on
rabbit populations varied but achieved an average reduction in rabbit numbers of 60% (Ramsey et al 2020).

An ‘RHD Accelerator’ approach was adopted to facilitate ‘in the lab’ accelerated natural selection of RHDV variants able
to overcome immunity to naturally circulating strains, or outperform them. This delivered proof-of-concept that virus
variants with altered immunological properties can be selected for and remain highly virulent in rabbits. However,
no variants were produced during this period mainly as a consequence of the need to carry out selection and growth
in experimentally infected animals. The approach has been paused until substantial progress has been made in the
development of culture systems that allow the cultivation of RHDV in vitro. This is being pursued through a joint
Meat & Livestock Australia and CSIRO Phase 3 project which aims to develop organoid tissue culture systems for
rabbits (see below).

Bioprospecting was chosen as the preferred method for a systematic review of known rabbit pathogens, with
assessments undertaken through expert consultations and stakeholders to determine their potential as suitable
biocontrol agents/biocides.

                         BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: AN UPDATE

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Current phase 3, 2017 to 2022
Phase 3 has continued to monitor the spread, evolution and interactions between the various virus strains in Australian wild
rabbit populations and is focussing on the case for RHDV2 to be registered as an additional tool for the rabbit biocontrol
pipeline. This involves undertaking a thorough characterisation of RHDV2 to assess its suitability as an additional biocide and
generating the data needed for an application to the APVMA for its registration.

A current CISS project is exploring the potential use of genetic biocontrol technologies as an alternative long-term, non-lethal
means of managing invasive animals (including rabbits) in Australia. These technologies often involve genetic modifications
to skew the sex bias of the offspring towards all-male, leading to the eventual collapse of the target population.

Meat & Livestock Australia and CSIRO are also co-funding a series of projects informed by and closely aligned with the rabbit
biocontrol pipeline to accelerate implementation of this phase of the pipeline strategy. Major project components include
the development of organoid tissue culture systems for rabbits and assessment of its suitability for the cultivation of rabbit
caliciviruses ex vivo; bioprospecting of lagomorph (rabbit and hare) pathogens within Australia and abroad; and modelling
using population genomics data to investigate whether emerging genetic control technologies may feasibly assist in the
control of rabbits in Australia.

Phase 4, 2022 to 2027
Based on consideration of the first three phases of the implementation of the long-term biocontrol R&D pipeline, this report
provides a framework and recommendations to move forward into Phase 4.

A strong proactive approach should be taken in Phase 4 that considers the available and potential short, medium and long-
term options, and balances the risks and likelihood of success with the potential benefits. The approach must maintain
critical capability and the ability to quickly react to new opportunities (pathogens or technologies) as they arise.

Phase 4 should highlight the importance of maintaining the underpinning scientific capability and infrastructure, as well as
the need to continue to improve integration of biocontrol applications with conventional control methods for maximum
impact.

                         BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: AN UPDATE

                                                              19
Rabbit biocontrol innovation pipeline.
 Achieving sustainable landscape scale rabbit management.

                                            PROBLEM                              SOLUTION

                       Numbers increase due to genetic
                        resistance to myxomatosis and           Post 1960s
                           weakening of myxoma virus.                         RHDV1 Czech strain released as
PRE-IA CRC

                                                                  1995        injection only ‘suspension’ formula.
                      RHDV1 Czech strain use expensive.                       ò 90% of rabbit populations (arid areas).
                     Needs to be frozen during transport.

                      RHDV1 Czech strain less effective in
                                                                  1998
                           cool-wet regions of Australia.

                                                                              RHDV1 Czech strain approved to mix
                                                                  2006        with baits. Allowing more efficient
                                                                              distribution of virus.

                                                                              Discovery of benign rabbit calicivirus
                                                                  2009        (RCV-A1). Identified as reason RHDV1 Czech
                                                                              strain less effective in cool-wet regions.

                                                                              Global search for new strain of RHDV to
                                                                              overcome benign strain.
                                                                              RHDV1 Korean strain (RHDV1 K5) selected.

                                                                              Assessment of potential rabbit biocontrol
                      Genetic resistance to RHDV1 Czech
                                                                  2015        agents which overcome genetic resistance
                             strain becoming a problem.
IA CRC

                                                                              issues and provide long term solutions.

                                                                              Strengthened National RHDV Monitoring
                       Outbreak of RHDV2 discovered in                        Program (incorporating RHDV2
                     Australia, full implications for RHDV1                   monitoring and online ability to report
                                K5 release not fully known.                   all evidence of dead rabbits).

                                                                              RHDV1 developed as a freeze dried
                                                                              formulation.
                                                                              Reduction to virus transport costs.

                Rabbits impacting 322 threatened species                      RHDV1 K5 proposed for release to overcome
                   in Australia and cause more than $200          2017        impact of native benign calicivirus. Available
                     million damage p/year to agriculture.                    as freeze-dried formulation.

                     Australians may get complacent that
                                                                              Phase 3 of the 20 year biocontrol
                  rabbits no longer need to be controlled
                                                                              research and innovation pipeline strategy.
                  and rabbit biocontrol is the silver bullet.

                                                                              Further R&D into the potential of RHDV2
                                                                  2018+       to become an additional registered biocide
                                                                              to maximise impacts. Specific vaccine
                                                                              development supported for non-target rabbits.
CISS

                                                                              Digital and face to face extension
                                                                              enhanced to ensure land managers have
                                                                              the best knowledge and tools available to
                                                                              manage rabbits into the future.

             Figure 9: Rabbit biocontrol innovation pipeline.

                                   BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: AN UPDATE

                                                                20
The self-disseminating nature of pathogenic biocontrols means that recurrent outbreaks are a significant advantage to rabbit
management at a landscape scale, particularly in remote areas. The suppressive effect of biocontrol agents provide has also
greatly improved the results of conventional control techniques such warren ripping, baiting and fumigation in a more cost
efficient and effective manner.

The benefits of rabbit biocontrol are not uniform across Australia (see Figure 10). RHDV1 has generally been more effective
in drier, lower rainfall areas and less effective in areas of higher rainfall. Rabbit abundance decreased on average by two-
thirds (67.3%) in low rainfall areas but only by a quarter (27.5%) in high rainfall areas (Neave 1999).

        Figure 10: Average decline in rabbit populations by rainfall zone after the arrival of RHDV1 (Neave 1999).

Given their wide distribution across Australia biocontrol is the most cost-effective way of managing rabbits and their impacts.

Environmental benefits
The recovery of native vegetation

Rabbit biocontrols have major environmental benefits for native plants, animals and ecosystems. A broad range of evidence
following the release of RHDV1 in 1995 provides greater understanding and insight into these benefits. It is likely that similar
benefits followed the release of MV in 1950, however, little data of this was collected at the time. However, the initial decline
in rabbit abundance from MV is comparable to RHDV1. As such it likely resulted in similar environmental benefits.

Reports of native vegetation recovery after MV are limited but include bladder saltbush (Atriplex vesicaria) (Hall et al.
1964), rangeland vegetation (Fenner and Ratcliffe 1965), and an extensive list of rangeland native plants including mulga
(Acacia aneura) and native pine (Callitris columellaris) (Lord 2000, quoting pastoralist Alan Bartholomaeus). The same, or
comparable, native plant species had significant post-RHDV1 recoveries, especially in semi-arid areas despite those regions
having below average rainfalls during those recovery periods (Sandell 2002, Murdoch 2005, Mutze et al 2008, Bird et al. 2012,
Mutze 2016).

                              BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: AN UPDATE

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Some case examples are illustrated below. Figure 11 highlights the distinct recovery of vegetation at Thackaringa Station,
Broken Hill between 2000 and 2012 using photographs taken from the same location. The recovery cannot be attributed
solely to RHDV1 as conventional rabbit control measures were also used. Significantly this degree of regeneration happened
despite sporadic periods of drought.

A reduction in rabbit abundance from biocontrol to below 0.5 rabbits per hectare allows native plant seedlings to survival
and grow in a way that is impossible to achieve with higher rabbit densities. Estimates of green photosynthetic vegetation
biomass were generated for the arid shrublands in northern South Australia after RHDV1 was established in 1995 using time
series Landsat satellite imaging and graphing the normalised difference vegetation index (NDVI). Figure 12 shows how NDVI
increased significantly across these shrublands after 1995 when rabbit abundance reduced by 95 per cent (Burrell et al.
2017).

       Figure 11: Rabbit control from RHDV1 and conventional measures led to significant regeneration of vegetation on Thackaringa Station,
       Broken Hill despite sporadic drought conditions (Photos: David Lord).

                            BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: AN UPDATE

                                                                          22
Landscape scale
                                                                          recovery of native
                                                                           vegetation with
                                                                             suppression

Figure 12: Recovery of arid shrublands in northern South Australia as detected by Landsat satellites (Burrell et al. 2017, Photo by David
Peacock).

                      BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: AN UPDATE

                                                                     23
Figure 13 shows the regeneration of native species at Turretfield, South Australia, in October 2017 as a direct result of the
rabbit density falling below 0.5 / ha post-RHDV2. Turretfield had been regularly monitored since 1996 (Bird et al. 2012).

            Figure 13: Regeneration of native species at Turretfield, South Australia, October 2017. post-RHDV2. Top left: Christmas bush,
            (Bursaria spinosa). Top right: Drooping sheoak (Allocasuarina verticillate). Bottom left: Drooping sheoak (A. verticillate). Bottom
            right: unidentified Acacia); (Photos: David Peacock).

                            BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: AN UPDATE

                                                                            24
Figure 14 shows how native orchid numbers increased once RHDV1 established in the Coorong National Park of South
Australia. An estimated 60 million native orchids were added to the 1,000-hectare creek area by 2002 (Bird et al. 2010).

       Figure 14: Increase in the native orchid population in the Coorong National Park following the establishment of RHDV1 and the suppression
       of rabbit numbers (Bird et al. 2010, Photo by Peter Bird).

                            BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: AN UPDATE

                                                                          25
Native mammal recovery
The management of rabbits at a landscape scale using biocontrol not only allows native plant species to recover, but also
aids the recovery of many rare and threatened native mammal species. For example, populations of spinifex hopping mice
(Notomys alexis) and plains mice (Pseudomys australis), common wombats (Vombatus ursinus), red kangaroos (Macropus
rufus) and western grey kangaroos (M. fuliginosus) all showed signs of recovery following the introduction of RHDV1 (Read
2003, Mutze et al. 2008, Bird et al. 2012).

Based on long-term monitoring, Pedler et al. (2016) concluded:

 “In arid inland Australia, the release of the rabbit biocontrol agent RHDV has been the single most important and cost-
 effective conservation action for small, threatened mammals (and a range of other taxa and ecosystems) in recent decades.

 The dusky hopping-mouse (Notomys fuscus), spinifex hopping mouse (Notomys alexis), plains mouse (Pseudomys
 australis), and the crest-tailed mulgara or ampurta (Dasycercus cristicauda) increased their abundance by up to 365
 per cent following the arrival of RHDV1 in the arid zone (Pedler et al. 2016).”

Figure 15 illustrates the recovery of threatened desert mammals at a landscape scale in north-east South Australia with
recorded detection and range size represented by coloured dots and shading. The orange dots are pre-RHDV1 release and
the blue dots are post-RHDV1 release.

           Figure 15: Landscape scale recovery of threatened desert mammals following the release of RHDV1. Map shows the north east 1/3 of
           South Australia. Yellow dots and yellow shading indicate records of detection and range size from 1970 to 1995. Dark blue dots and shading
           indicate expansion of records of detection and range from 1996 to 2010. Light blue dots and shading indicate expansion of records of
           detection and range from 2010 to 2014. (Pedler et al. 2016, Photo by Reece Pedler)

                           BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: AN UPDATE

                                                                         26
The recovery of these native animals is intrinsically linked to the recovery of their habitat and food resources following the
reduction in rabbit abundance. An additional benefit further enhancing their recovery was the reduction in the abundance of
foxes (Vulpes vulpes) and feral cats (Felis catus) that was directly attributed to the removal in their primary prey animal, the
rabbit (Cooke & Soriquer 2016).

Fox and feral cat numbers were shown to have decreased in the Ikara-Flinders Ranges National Park (South Australia) after
the rabbit population had been reduced by 85 per cent following the introduction of RHDV1 (Holden & Mutze 2002). A
similar decline in feral cat populations was also reported at Roxby Downs in South Australia after the arrival of RHDV1 (Read
and Bowen 2001) with evidence of increased hunger and reduced cat survival following the significant reduction in rabbit
abundance (McGregor et al. 2020). This impact is illustrated in Figure 16.

        Figure 16: The landscape scale suppression of rabbits, foxes and feral cats in 1989–90 at Roxby Downs, South Australia, following the arrival
        of RHDV1 (Read and Bowen 2001; Photo by Scott Jennings).

                             BENEFITS OF RABBIT BIOCONTROL IN AUSTRALIA: AN UPDATE

                                                                            27
Although the data is limited, it is likely that declines in feral cats and foxes, together with subsequent benefits to native fauna
also occurred following the introduction of MV which saw rabbit abundance decline by up to 99 per cent across Australia.

There is a clear link between the prevalence of mange and nutritional stress in many animals including wombats (V. ursinus)
(Skerratt et al. 1998) and other species (Pence & Ueckermann 2002). In 1952, an Australia-wide increase in the incidence of
manage in foxes led to a major decline in their condition and abundance (Rolls 1969). It is likely that the sharp decline in
rabbit numbers that followed the release of MV in 1950 led to this outbreak of mange in the fox population.

Bottom-up rabbit trophic cascades — where animal population abundance is governed by available food and nutrition —
are important drivers of the sustainability and recovery of the Australian environment. Widespread suppression of rabbits
not only enables the direct recovery of susceptible native vegetation and the fauna that rely on it, but it also benefits native
fauna through increased habitat and food resources and the suppression of feral cat and fox abundance.

Carbon benefits through biosequestration
Biosequestration refers to the capture and storage of the atmospheric carbon dioxide, largely through increased rates of
photosynthesis resulting from increased vegetation biomass and revegetation. There is a strong temporal relationship
between ground cover and vegetation biomass and soil organic carbon levels for given climatic conditions and soil types.

The introduction of RHDV1 in 1995–96 saw declines in rabbit populations of up to 98 per cent (Mutze et al. 1998) with a
corresponding increase in standing biomass observed by satellites (Burrell et al. 2017). Therefore, continued investment in
biocontrol agents to reduce rabbit numbers is likely to provide environmental benefits resulting from an increase standing
vegetation biomass and, consequentially, carbon sequestration (Hardaker & Chudleigh 2020).

Economic benefits
Prior to the release of the myxoma virus, the financial impact of rabbits on agriculture was some $2 billion a year. Rabbit
biocontrol based on the myxoma virus and RHDV1 Czech 351 strain reduced this impact to about 15% of the pre-release
cost (Cox et al 2013). Based on their review of the available literature and the application of a loss-expenditure frontier
model with and without biocontrol scenarios, Cooke et al. (2013) conservatively estimated the cumulative benefit of MV and
RHDV1 to Australia’s livestock and farming industries at approximately $70 billion from 1950 to 2011. This value is expressed
in 2011-dollar terms and is made up of around $54 billion from MV on its own and an additional $16 billion from MV and
RHDV1 together. Expressed in 2020-dollar terms, this equates to a cumulative benefit of $81.8 billion over
six decades(Reserve Bank of Australia, 2022).

The Cooke et al. analysis was undertaken prior to two further major events that have impacted rabbit populations. In
May 2015 a new variant of RHDV was detected in New South Wales and serological studies showed it had been present in
Australia since at least the Spring of 2014. It was a highly virulent RHD virus, known as RHDV2, that spread rapidly across all
Australian states and territories within two years of its arrival. It was estimated to have reduced average rabbit abundance by
approximately 60 per cent (Ramsey et al 2020).

The other development post-2011 was the 2017 release of the K5 strain of RHDV1. The appearance of RHDV2 in Australia
has been shown to have severely suppressed the potential benefits of the release of RHDV1-K5 (Ramsey et al 2019) and the
Centre for Invasive Species Solutions commissioned Hardaker and Chudleigh (2020) to evaluate the impact of RHDV1-K5 for
control of rabbits in Australia. The evaluation aimed to estimate:

      — the projected potential impact of RHDV1-K5 (without the emergence of RHDV2), based on available data for
        RHDV1-K5 and RHDV2 impacts; and

      — the current impact of RHDV1-Czech-351, RHDV1-K5 and RHDV2 together, and then assess the actual impact
        attributable to RHDV1-K5.

Hardaker and Chudleigh’s economic analysis calculated the estimated Present Value of Benefits for RHDV1-K5 over 30 years
using a 5 per cent discount rate (see Table 2). Their analysis produced the following results:

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